Release and establishment of Harmonia axyridis (Coleoptera: Coccinellidae) in the south- eastern United State

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228 ENTOMOLOGICAL NEWS

RELEASE AND ESTABLISHMENT OF

HARMONIA AXYRIDIS

(COLEOPTERA: COCCINELLIDAE)

IN THE SOUTHEASTERN UNITED STATES 1

W. L. Tedders 2 , Paul W. Schacfcr 3

ABSTRACT: Harmonia axyridis were laboratory reared and 87,810 specimens were released
{1978-8 1 ) primarily in pecan orchards containing leguminous cover crops at Byron, Georgia, The
first major recovery indicating establishment in Georgia was in 1990 at Buchanan, 174 km from
the Byron release site. By 1992, the population had spread throughout Georgia to northern Florida
and eastern South Carolina. H axyridis is now the dominant coccinellid species {54,4%) in the
Byron pecan orchard and appears influenced by the legume cover crop. Numerous pecan growers
in Georgia reported control of pecan aphids (Monellia caryella and Monelliopsis pecunis) dunng
1993. Adult H axyridis were more attracted to white traps than to similar dark grey, brown, or
black traps During rearing, total development required ca. 36 d. Recorded prey include 36 species
of Aphididae, 12 species from 7 other homopteran families, and 3 species from two other insect
orders. Aphids appear to be preferred prey but all may be acceptable, indicating polyphagy and
suggesting an extended control potential for this generally arboreal coccinellid

Lady beetles in the genus Harmonia occur in Asia and Australia and prey
on aphids, psyllids, and scale insects (Gordon 1985). Widely distributed in
Asia, Harmonia axyridis (Pallas) is recorded from Formosa, China, Korea,
Japan, Manchuria, southern Siberia and the Ryukyu and Bonin Islands
(Chapin and Brou 1991). This arboreal species occurs in orchard and forest
habitats and preys mostly on various aphids, but it will also accept certain
scales and two species of chrysomelids. It inhabits various trees, including
maple, walnut, willow, and oak {Chapin and Brou 1991). Harmonia axyridis
is highly polymorphic, very prolific, and may live up to Lhrec years (Hodek
1973).

Several attempts have been made to introduce H. axyridis into North
America, Inclusive dales for recorded releases include: California - 1916,
1964 and 1965; Connecticut, Delaware, Georgia, Louisiana, Maine, Maryland,
Mississippi, Nova Scotia, Ohio, Pennsylvania, Washington, and Washington,
D.C - 1978 lo 1982; and Connecticut - 1985 (Coulson el al. 1981, Gordon
1985, McCIure 1987; USDA, Beneficial Introduction Res. Lab., unpubk
records),

Recently, Chapin and Brou (1991) reported the establishment off/, axy-
ridis in Louisiana and Mississippi. Releases in that area consisted of a single

' Received February 4, 1994. Accepted February 26, 1994,

2 Southeastern Fruit and Tree Nut Research Laboratory, 1 1 1 Dunbar Road, Byron, Georgia
31008.

3 Beneficial Insects Introduction Research, 501 Chapel Street. Newark, Delaware 19713.

ENT. NEWS 105(4); 228-243, September & Ociober 1994

Vol. 105, No. 4, September & October, 1994 229

release of 32 specimens in Bossier Parish, Louisiana, in 1979 and eight
releases totaling 3781 specimens in Washington County, Mississippi (near
Leland), in 1980. The first evidence of establishment was the collection of
more than 1000 adults in six light traps near Abita Springs, Louisiana, from
July to November 1988. Specimens were collected from August to November
1990 in the Mississippi counties of Calhoun, Harrison, and Panola, In addi-
tion, Chapin and Brou reported specimens from crabapplc trees infested with
Aphis spiraecola Patch, crapernyrtle infested with Tmocallis kahawaluokalani
(Kirkaldy), and from senescent Erigeron sp. in an old field habitat. Currently,
K axyridis is the most common lady beetle in the Abita Springs area.

Ill is paper records the appearance of H, axyridis in Alabama, Georgia,
South Carolina, and Florida. We also report on the rearing and release pro-
gram in Georgia (Gordon and Vandenberg 1991, Tedders 1986, Tedders 1991)
and give additional notes and information concerning its biology and behavior
in Georgia and its known prey.

MATERIALS AND METHODS

Rearing and Production. An intensive H. axyridis rearing and release
program was conducted at the USDA-ARS, Southeastern Fruit and Tree Nut
Research Laboratory at Byron, Georgia, from 1978 to 198L The target prey
were black pecan aphid, MelanocaUis caryaefoliae (Davis), blackmargined
aphid, Monellia caryella (Fitch), and yellow pecan aphid, Monelliopsis
pecanis Bissell, on pecan, C&rya illinoensis (Wangenh.) K. Koch. Stock
insects were supplied by the USDA-ARS Beneficial Insects Introduction
Research Laboratory at Newark, Delaware. At Byron, most H. axyridis were
reared on Myzus persicae (Sulzer) colonies that infested greenhouse-produced
Chinese cabbage, Brassica rapa L. (pekinensis group), Chinese cabbage
plants were grown in Jiffy-9® peat pellets, Additional fertilizer, fungicides,
and insecticides were not used. Two tightly secured greenhouses were used,
one to produce aphid-free cabbage plants and the second to produce cabbage
plants infested with M. persicae. Non-infested plants were transferred to the
second greenhouse where they remained for a few days and became infested,
Aphid-infested plants were then removed and placed in individual cages con-
taining one or more pairs of H. axyridis adults.

Cages for ft axyridis were 3.8 £ cylindrical cardboard cartons having a
thin polyethylene upper cover that allowed for illumination and viewing.
Newly infested cabbage plants were added to each cage every second day or
as needed as aphids were consumed. Cabbage plants were provided in cages
at the rate of one plant per pair of H. axyridis. Cabbage plants depleted of
aphids were discarded. The inside of the carton cages were lined with brown
wrapping paper as an oviposition substrate. Foliage, the paper liner, and the

230 ENTOMOLOGICAL NEWS

cover were removed daily to obtain eggs for additional colony production for
release, developmental studies, and studies of acceptance of pecan aphids as
prey. Specimens of stock lady beetles were maintained in the insect collection
at Byron as identification vouchers.

In the greenhouses, temperatures averaged 24.6°C (range 6.7 - 37.7°C) and
relative humidity averaged 68.5% (range 30 - 100%). In the rearing facility the
temperature averaged 23,6°C (range 20 - 27.7°C) and relative humidity aver-
aged 69.4% (range 40 - 95%). Lighting was held constant by overhead lights
24 hr per day. The production cages often contained 2 or 3 pairs, and supply
of food was increased at a comparable rate.

In November 1992, visual traps developed for monitoring the emergence
of pecan weevils, Curculio caryae (Horn) (Stanley 1992, Tedders and Wood
1993) were used to assess visual color preferences. The traps were interlock-
ing triangular masonite panels, measuring 53.3 cm base x 121.8 cm height, on
top of which was positioned a modified boll weevil, Anthonomus grandis
grandis Boheman, collecting device (Anonymous 1990). The panels were
painted either white (84.0% reflectance), gray (18.3% reflectance), brown
(4.8% reflectance), or black (1.0% reflectance). Each colored trap was repli-
cated three times and set out across the lawn of the laboratory in a randomized
complete block design. Traps were about 3 m apart. Traps were in place from
about 3:00 p.m. November 17 until 3:00 p.m. November 20 (3 consecutive 24-
hr periods). Beetles were removed and counted after each 24-hr period. Trap
data were subjected to analysis of variance.

RESULTS AND DISCUSSION

Rearing and Development- Eggs were laid on the paper lining, cabbage
foliage, and the polyethylene cover. A shortage of aphid prey in the cages was
always followed by a marked reduction in lady beetle eggs due to cannibalism.
Well-fed beetles did not usually cannibalize their own eggs.

Eggs averaged 4 days from oviposition to hatch (range 3-5 d). Average lar-
val development for first through fourth stadia was 2, 2, 2-4, and 6 d, respec-
tively, with larval development overall averaging 12-14 d, The pupal stadium
ranged from 5-6 d and averaged 5.5 d. Average time required for development
from egg to egg was about 36 d. Cooler temperatures ranging from 15.5 to
21J°C increased the time required for development from egg to egg by 3-4 d.

Thirty-one newly emerged females were studied for longevity and fecun-
dity. With one copulating pair confined per cage, females lived an average of
32.2 d (sd=25.26), laying an average of 491.3 eggs (sd=375.86); mean 16.7
eggs/female/d (sd=7.64). The shortest life span was 2 d and the longest 101 d.
The 1 01 -day old female produced 1543 eggs. Longevity of males was not
recorded hut was usually shorter than that of the females.

Vol. 105, No. 4. September & October, 1994 231

Adults and larval mortality in all colonies was usually the result of natural
causes such as individual weakness and old age. Dead beetles were placed in
a humidor and observed for suspicious microbial growth. On two occasions
fungal pathogens were suspected and identified as Paecilomyces farinosus
(Holm) Brown and Smith (Deuteromycota) (Richard A. Humber, pcrs. comm.,
Boyce Thompson Institute, Ithaca, New York).

All three species, M. caryaefaliae, M caryella, and M. pecanis represented
adequate food when time required for larval development and oviposition on
a diet of M. persicae were used as baseline data. However, H. axyridis
appeared to prefer Atf, caryella. The preference may be associated with the
copious honeydew excreted by this aphid because both larvae and adults used
this alternative food source. Adult H. axyridis, fed on M. pecanis or M.
caryella, laid about 20 eggs per day. Melanocalhs caryaefoliae, which
appeared to be least favored as prey, excretes the least amount of honeydew.

Releases. Releases of reared H. axyridis in Georgia from 1978 to 1981
usually consisted of second and third instar larvae, occasionally 1st instar lar-
vae, and rarely eggs. In total 87,561 immatures and 249 adults were released
from 1978 to 1981 (Table 1). Releases were made on the farm at the Byron,
Georgia laboratory, mainly into trees or legume cover crops within pecan
orchards. Immatures were either transferred to plant foliage with a camel-hair
brush, or the supporting substrate of Chinese cabbage leaf, paper liner, or
polyethylene cage cover was stapled to pecan or legume foliage. Pecan trees
were selected that supported Af caryella, M. pecanis, or M, caryaefoliae
aphids or mixtures of two or all three species. The primary release orchard
(about 7 ha) was bordered on opposite sides by oak-hickory-gum woods, on a
third side by a second pecan orchard, and on the remaining side by a fallow
field. No pesticides were applied to the orchard from 1977 through 1983,

Releases during March, April, and May were usually divided evenly
between winter cover crop in the orchard and the pecan trees. The winter cover
was composed of hairy vetch, Vicia villosa Roth, crimson clover, Trifolium
incarnation Roth, big flower vetch, Vicia grandiflora Scopoli, and rye grass,
Loliitm sp. Vetch and clover ground cover infested with pea aphids,
Acyrthosiphon pisum (Harris), and occasionally cowpea aphids, Aphis cracci-
vora, and A, medicagenis Koch was selected. Larvae and adults (n = 397)
were released in nearby cultivated plums, Primus sp., apple, Mains sp., and
crapcmyrtle, Lagersiroemia sp. Plums were infested with black peach aphids,
Brachyc audits persicae (Passerini), rusty plum aphid, Uysleroncura setariae
(Thomas), or A. spiraecola. Apple was infested with A. spiraccola and crape-
myrtle with T. kahawaluokakmL Also, a total of 2,485 larvae were released in
weed covers within pecan orchards where I he dominant weeds were horse-
weed, Erigeron canadensis L., infested with Uroleucon erigeronensis
(Thomas), cocklebur, Xanthium strumarium L., infested with I '. utnhmsiae
(Thomas), and goldenrod, Solidago sp. infested with LL rudheckiae (Filch)
(Table 1).

232 ENTOMOLOGICAL NEWS

Initial Recoveries. From 1978 to 1981 attempts to recover adults of
released H. axyridis were conducted weekly, beginning the first week of April
and continuing until the end of October or mid-November depending upon the
condition of pecan foliage and the weather. From 1982 to the present, collec-
tions of beneficial arthropods at Byron, emphasizing Coccinellidae and
Chrysopidae, continued on the same schedule in the same 7-ha release orchard
or in adjacent orchards not over 8 km distant. Given these efforts, if coccinel-
lid establishment had been successful, beetles should have been detected.

From 1978 to 1981, three sampling methods were employed in the 7-ha
orchard. Visual 2-hr searches were conducted weekly in and around the
orchard. In addition, weekly sweep net samples of the ground cover (100
sweeps per 7 ha) were taken, and the lower limbs of 25 pecan trees within the
orchard were jarred with a club to dislodge beetles onto a 1 m 2 cloth catching
frame.

From 1982 to present, visual searches of about 1 hr were conducted
monthly. Sweep net collections from cover crops continued but collections
from trees were made with the aid of a D-Vac collecting machine as a substi-
tute for the limb-jarring method.

After releases, mature larvae and pupae were easily found on foliage of
pecan trees and the ground cover, however adults were scarce. One adult was
swept from evening primrose, Oenothera sp.. May 24, 1978. No adults were
collected during 1979. Six adults were swept from vetch from April 28 to May
6, 1980, and fifty- three adults were collected from pecan foliage by jarring and
by hand collection from May 5 to June 6, 1980. One adult was swept from rye
grass on May 6, 1980, A single adult was recovered September 29, 1981, by
jarring pecan foliage. No adults or larvae were collected from 1982 through
1991 .

Evidence of colonization in Georgia first became available in fall 1990.
John C, Callaway, Jr., County Extension Director for Haralson and Carroll
Counties (Buchanan, Georgia) advised the Department of Entomology, Uni-
versity of Georgia, Athens that an unidentified lady beetle had become a nui-
sance in houses in that area. A visit there on June 18, 1991 revealed a single
H, axyridis adult on arrow leaf clover, Trifolium vesiculosum Savi, No addi-
tional lady beetles were found on adjacent trees, shrubs and other vegetation,
but suspect pupal exuviae were found on apple. Specimens of that population
submitted for identification to the Department of Entomology, University of
Georgia, proved to be H, axyridis (Cecil L. Smith, per. comm).

A second trip to Haralson County (elevation 382 m) on November 12,
1991, revealed about 50 H. axyridis adults inside the screened porch of a home
in Buchanan. Additional searches revealed numerous adults and pupae on
apple, Finns spp., and magnolia, Magnolia macrophylla Michaux, at this site.
During November and December 1991 Mr. Callaway and the Byron Labora-
tory received numerous reports that H. axyridis was an increasing nuisance in

Vol. 105, No, 4, September & October, 1994 233

homes throughout northwest Georgia. During early winter of 199 1 H. axyridis
were identified from northeast Alabama and were abundant near Huntsvitle
(Paul Estes, Auburn, Alabama, per. comm.).

Subsequent Recoveries-Biology and Population Abundance, At Byron,

on February 4, 1992, one adult was collected from loblolly pine, P. taeda L.
that was heavily infested with Eulachnus agilis (Kal ten bach) on which the
beetle was feeding. Thereafter and throughout March adults, larvae, and eggs
were found regularly on various species of yellow pine infested with E. agilis.
Foliage samples from April through October 1992 revealed that H, axyridis
began inhabiting pecan in May and fluctuated in numbers as aphid populations
varied, Harmonia axyridis comprised 54.4% of all the lady beetles collected
from pecan by the end of the growing season (Table 2).

In a separate experiment on October 6, five terminal branches on each of
20 pecan trees were examined for H. axyridis by the jarring technique. Adults
averaged 0.30/terminal and larvae 0.27/terminaL Aphid control on pecan,
which was considered to be very good throughout 1992, was attributed mainly
to H. axyridis ,

By late summer 1992, specimens of H, axyridis were found throughout
Georgia (W. L. T.), in north Florida (Russ MizelK University of Florida, pers.
comm.) and near McClellanville, South Carolina (Stuart H. Tedders, Univer-
sity of South Carolina, pers. comm.).

Movement by M axyridis to overwintering aggregation sites began about
November 1, 1992, when several hundred were found on the southwest sides
of several buildings at Byron. On November 17 adults were observed flying
around the grounds of the laboratory in such abundance that they were first
thought to be the tail end of a swarm of honey bees. Many of these landed on
the sides of buildings in large numbers, and four people were able to hand col-
lect about 6,000 beetles in 2V 2 hrs.

Since beetles appeared to be attracted in greater numbers to light colored
buildings than to dark colored ones, it was not surprising that traps captured
totals of 208, 143, 61, 23 in the white, gray, black, and brown traps, respec-
tively. Analysis of variance of these data indicated that H. axyridis is signifi-
cantly attracted to white traps (a = 0.05) as opposed to darker ones (Table 3).
This finding confirms observations by Obata (1986) in Kyoto, Japan, and
agrees with those of Hodek (1973) that several coccinellid species were
attracted to light-colored rock outcroppings that were used as overwintering
sites. Beetles were not observed moving to overwintering sites after Novem-
ber 20, 1992.

Recoveries began in 1993 when several adults were found (February 15)
feeding on the floral nectaries of common box, Buxus sempenirens L„ indi-
cating that emergence from overwintering had begun. About 2,000 hibernat-
ing adults were collected on February 26 by three people in 30 minutes from
the center of a panelled wall on the south-southwestern side of an abandoned

234 ENTOMOLOGICAL NEWS

insecticide building at the laboratory farm center, indicating that most were
still overwintering. Also on that date, several active adults were observed on
yellow pine, and about 30 active adults were observed on the lighting fixtures
in an office of the laboratory building, further indicating that the overwinter-
ing period was ending.

At Biloxi, Mississippi (February 28), mature larvae, pupae, and newly
emerged adults were found on Podocarpus sp, infested with Neophyllaphis
podocarpi Takahashi at a hotel resort facing the Gulf of Mexico, confirming
that H. axyridis had been active in that area for several weeks, and had passed
through at least one generation (W.L.T.).

On March 9 at Byron, adults were observed feeding on exudates of the flo-
ral nectaries of peach, plum, and common box. Aphids or other prey were not
observed on these plants, and we can exclude the possibility that aphid eggs
were being fed on since aphids are rare on these plants in central Georgia. On
March 1 1 7 100 limbs each of peach, plum, and yellow pine were jarred reveal-
ing 13, 1, and 3 adults, respectively.

Impact in Pecan orchard: Collections of//, axyridis from pecan orchards
at Byron continued during 1993. In addition to standard sweep net and D-Vac
machine samples, a large Malaise trap was placed in a legume ground cover
study orchard. Collections from this orchard were compared, with collections
from an adjacent control orchard without a legume cover. Yellow aphids on
pecan in the legume cover crop orchard were very low, the highest count aver-
aged only 4,7/compound leaf during the week of May 10. Average numbers of
aphids for May 1993 was 2.2 aphids per compound leaf. Yellow pecan aphids
on pecan during May usually exceed 100/leaf. In the adjacent control orchard
the highest counts were 22.3 aphids per compound leaf during the week of
May 17, and counts averaged 9.2 aphids per compound leaf for the month.
Neither orchard received insecticide but H. axyridis were noticeably slower to
colonize the non-cover crop orchard. Although a Malaise trap was not used
and sweep net samples were not taken from the closely mowed grass of the
control orchard, comparative collections of//, axyridis suggest the importance
of the legume cover crop (Tabic 4). Numerous pecan growers in Georgia
reported excellent control of M. caryella and A/, pecan is in commercial
orchards during 1993 (W.L.T.).

Conclusions: Harmonia axyridis, first reported to be established in North
America by Chapin and Brou (1991) from Louisiana in 1988 with subsequent
collections during 1989 and 1990, was not reported in Mississippi until 1990.
There was no mention by the authors of the size of that population. Releases
possibly leading to the Louisiana-Mississippi establishment were a single
release of 32 specimens in Bossier Parish during August 1979 and eight
releases of 3781 specimens from July to October 1980 near Leland, Missis-
sippi. The straight-line distance from Bossier City (Bossier Parish) to Abita

Vol. 1G5, No, 4, September & October, 1994 235

Springs, Louisiana is about 418 km (260 mi) and from Leland to Abita Springs
is 320 km (199 mi), The distance from Leland and Bossier City to Buchanan,
Georgia is about 553 km (344 mi} and 83 1 km (516 mi) respectively, and from
Abita Springs to Buchanan is about 591 km (367 mi) (Figure 1).

The population level first reported in Georgia near Buchanan during the
fall of 1990 was very high as judged by numerous calls to county agent John
Callaway in Buchanan (personal communication). The straight-line distance
from Byron to Buchanan is about 174 km (108 mi). It remains uncertain
whether the Buchanan population was the result of migration of beetles across
Mississippi and Alabama to Buchanan or the result of migration of releases
from Byron to Buchanan. It is clear that many more specimens were released
at Byron and the distance from Byron to Buchanan is much shorter. If the
Buchanan population originated from Abita Springs, it is unusual that H.
axyrdis were not reported as a nuisiancc from Alabama before it was found in
Georgia in view of its habit of overwintering in homes. The evidence suggests
that two separate establishments occurred: one in Abita Springs, LA, and
another in Buchanan, GA.

Because H. axyridis is highly polymorphic, with a basic red morph and a
black morph, as illustrated in Ayala (other variations are also possible), it is
important to record that no black morphs were found in Georgia, Alabama,
Florida, and South Carolina populations. All beetles found to date are of the
red color morphs as described by Chapin and Brou (1991 ). Live specimens of
red morph vary in background color from a pale yellow-orange to a deep
orange-red (which may well be age related), and the black spotting varies from
none to 20± spots.

As //. axyridis spreads from the southeast into other areas and begins to
receive increasing attention, a greater understanding of this invading species
will result. Toward that end, we present a rapidly growing list of prey species
(Table 5), This tentative list will serve as a reference point for addition of new
prey species, especially as this lady beetle is redistributed (See Appendix) and
spreads naturally into new states,

APPENDIX

In light of some significant biological information and records of an inten-
tional shipment of Harmon ia axyridis, we offer the following information for

the record.

Biological Notes; April 1 6, 1 980, freeze-dried pea aphids and a water sup-
ply were found to be acceptable but poor quality food for //. axyridis . Thirty-
six eggs yielded five pupae but only three adults.

\9&0-Uroleucon amhrosiae on horseweed and coeklcbur were unsatisfac-
tory prey and H. axyridis did not complete development, Uraleucon erigeron-
sis on horseweed were fair prey and produced adults after lengthy feeding.

236 ENTOMOLOGICAL NEWS

Aphis craccivora on vetch and velvet bean, Stizo labium Deeringianum Bert.,
were poisonous to H. axyridis larvae. Aphis spiraecola on Garland cv. crab
apple, Mains coronaria Mill, were poor prey for larvae.

May 22, \98\-Phylloxera notabilis Pergande on pecan were excellent
prey for H. axyridis; larvae matured in less that 12 d. Field released adult
H. axyridis were observed feeding on P. notabilis emerging from opening galls.

August 25, \9$\-Tinocallis kahawaluokalani on erapemyrtle were suit-
able prey. Larvae released on erapemyrtle yielded at least two adult H, axy-
ridis , Four H. axyridis adults fed A. pisum laid a total of 2,016 eggs; average
526.5 eggs/female over a period of 32.3 consecutive days. As prey, A. pisum
were deemed suitable.

For initial rearing of H. axyridis in quarantine at Newark, prey were A.
pisum produced on faba (fava) beans, Viciafaba L. Cages for all life stages
were 530 ec unwaxed paper cups (No. 2186 Design, James River Corp.) with
tight-fitting clear plastic lids. To prolong the availability of suitable A. pisum
prey, several lengths of bean stems were stripped of leaves and placed in each
cup. By stripping stems of leaves, the containers were kept cleaner, which
facilitated detection and removal of eggs, larvae, and adults.

Samples of overwintering //, axyridis were weighed on two occasions after
collection to gain information about weight loss during hibernation. On
December 15, 1992, 315 beetles weighed an average of 35 mg. On January 1 1,
1993, 224 beetles weighed an average of 33 mg for loss of 2 mg over 27-d
period.

May 1993— All stadia of H. axyridis were found in very large numbers on
six tulip trees, Liriodendron tulipifera L., infested with Macrosiphum lirio-
dendri (Monell) in Houston County, Georgia.

April 199 3 -Ornamental rose, Rosa spp., infested with Rhadobium poro-
sum (Sanderson) were found to have one or more feeding adult or larval //,
axy'ridis per plant in Thomas ville, Ga, The H. axyridis population was very
large and effectively controlled aphids during this time.

Redistribution: Of the 8,000 H, axyridis adults collected during late 1992
and early 1993, 6,000 were sent to California for inspection and subsequent
release in California and New Mexico pecan orchards (Ken S, Hagen, Kent
Daane, and Steve Sibbett, University of California, pcrs. comni). To date
1,500 were released on April 9 and 750 were released on April 30 at Blaine
Ranch, Visalia, California. Dr. Hagan inspected and forwarded 2,000 to New
Mexico where they were released in pecan orchards at Mesilla (Joe Ellington,
New Mexico State University, pers, comin)

During fall 1993, daily observations were made to detect the onset of H.
axyridis flight to overwintering quarters. None were observed until November
4 when large numbers congregated on the sides of buildings at Byron (maxi-
mum air temperature was 22.2° C). Very few were observed on November 5

Vol. 105. No. 4, September & October, 1994

237

(maximum temperature 20.6° C) and fewer yet occurred during the next four
days when maximum temperatures were 16.6° C or below.

Maximum flight activity occurred on November 10 when many adults
congregated on the sides of buildings. Of particular interest that day was an
abandoned silo located on the laboratory farm at Byron. The silo was a light
gray concrete shell having no top and containing about 20 cm rain water at the
bottom. Insects on the inside were inaccessible but visible through port holes.
The silo measured 14.4 m cir. x 9.6 m height, At about 2:00 p.m., congregat-
ing //. axyridis were estimated at 1550 beetles/m2 on the outside wall with
slightly smaller numbers on the inside wall. Air temperature for November 10
was 20° C maximum and 5° C minimum.

On November 1 1, entomologists Billy Ree and Allan Knutson of the Texas
Agricultural Extension Service, Texas A&M University, arrived at Byron to
collect H. axyridis during November 11-14. They collected 10,200 adults,
mostly from the exterior wall of the silo. Maximum temperature for the period
ranged from 19.4-25,0° C. Smaller numbers of adults were observed on the

Boaaler

Figure I. Map of Southeastern United States showing release sites (Byron, GA. Ldand MS,
and Bossier City, LA) and recovery sites (Buchanan, GA and Ahita Springs LA) of Harmoma
axyridis and the straight-line distances (KM) between sites.

238

ENTOMOLOGICAL NEWS

silo on each successive day, but the change was not measured. On each day,
the primary movement of adults to the silo occurred between 1:00 and 4:00
p.m. when air temperature ranged between 17.7-20.6° C. During that period,
the sky was bright but large cumulus clouds intermittently blocked full sun.
The greatest number of adults was observed flying to the silo during periods
when the silo was fully illuminated, and flying decreased with shadows cast
by clouds. Most H. axyridis that accumulated during the day were gone the
following morning; presumably because of a lack of protected sites (cracks
and crevices) to inhabit. The H. axyridis collection was returned to Texas for
study and release in Texas pecan orchards during spring 1994. One Byron
technician collected 2795 H. axyridis from the silo for our use on November
12 during a 1.5 hr. period. Adult weight averaged 35.4 mg.

Table 1 Release of Hurmoma axyridis into pecan trees and clover-vetch ground cover, Byron,
Georgia.

1978

1979

1980

1981

Imrnatures

Adults

Imrnatures

Adults

Imrnatures Imrnatures

Mar.

23,401

Apr.
May
Jun

197

1.195

2,496

34

162 2

6

1.767

723

18

6.729

1 ,052

Jul

482

Aug
Sep
Oct
Nov

258

2,780

2,846

23

12,164
2.145 4

15,244
12.318 5

933

809

22

Total

10.254

225

44,361

24

3 1 .204

1,742

I

201-plum, 172 apple
£ 18-apple
■ 6-crapemyrtle

4 540-weed cover
-* 1,945-weed cover

Vol. 105, No. 4, September & October 1994 239

Table 2. D-Vac collections of important Coccinellidae from pecan trees at Byron, Georgia, 1992.

Species MonthyNumber Collected

Apr May Jun Jul Aug Sep Oct Total

Hippodamia converge ns 422 000 8

Coccinella sepiempunctata 2 16 8 5 I 2 34

Olla v-nigrum I 4 5 4 16 2 1 33

Cychmeda munda

IY

Coteomegilta maculata 1 4 11 16

Harmonia axyndis 2 9 53 34 34 132

Total 6 32 IS 27 73 39 47 242

Table 3. Analysis of variance of means of numbers of Harmonia axyridis adults captured in four
different colored traps November 17-20, 1992, Byron, Georgia 1 .

Trap 24 hr periods^ Total of 3

Color 1 2 3 Penod 3

White 5.4 a 5.6 a 2.6 a 8.1a

Gray 4.3 ab 4.2 a 2.1 ab 6.6 ab

Black 2.8 be 3.0 ab 17 ah 4.5 be

Brown 2,0 c 1..3b 0.7 b 2.8 c

MSE 0.899 1767 0.592 2,164

' Analysis of variance of means; a = 0.05, df - 6. Means followed by same letter within a col
umn are not significantly different (Duncan's Multiple Range Test).

- Means captured in 3 traps per 24 hi period

-^ Means captured in 3 traps per 72 hr penod.

240 ENTOMOLOGICAL NEWS

Table 4. Marmonia axyridis captured in pecan orchard with ground cover of crimson clover and
hairy vetch, sampled by malaise trap, sweep net, and D-Vac methods, compared with adjacent
orchard with no cover crop, sampled by D-Vac only, Byron, Georgia 1993.

Legume Cover Crop Orchard No Cover Crop

Total 29 51 15

Sweep net samples represent 5 sweeps in each of 10 locations.
* D-Vac samples represent 5 terminals/tree from each of 10 trees

Sample Malaise Sweep

Period Trap Net D-Vac- D-Vac 6

Mar 8-14 — 2 — —

Mar 15-21 1 — —

Mar 22-28 2 2 — —

Mar29-Apr4 8 10 — —

Apr 5-11 4 23 — —

Apr 12-18 8 3 3

Apr 19-25 6 5 2 1

Apr26-May2 3 4 1

May 3-9 10 11

May 10-16 2 1

May 17-23 10 1

May 24-30 I 3

Vol. 105, No, 4, September & October, 1994 241

Table 5, Recorded prey of Harmonia axyridis Pallas.
Taxon and Species [Source(s)] 1

Coleoptera:
Chrysomelidae

Amhmstoma quadriimpressum Motschulsky 4
Chrysomela vigintipuncaia Scopoli
Homoptera;
Adelgidae

Adelges laricis Vallot
Aphididae
Acyrthosiphon pisum (Harris) 2 ' 3 (Hodek 1973)

Agrioaphis spp.

Amphowphora oleracea v.d. Goot J

Aphis craccivora Koch (Hodek 1973)

Aphis ponti DeGeer (Hodek 1973)

Aphis spiraecola Patehr (Chapin & Brou 1991 )

Cfkiiiophorus spp.

Cinara kochi Inouye

Cinara laricicola (Matsumura) 3

Cinara lodocola (Inouye) 3

Cinara pinea Mordvilko 4

Cryptosiphum gallarum Kaltenbach-*

Eriosoma lanigerum (Hausirannr

Eulachnus agilis ( Kaltenbach) (Present study)

Hvalopterus pinni Matsumura

Hyaiopterus pruni (Geoffrey) 3

Kermaphis pini (Koch)^

Lachnus sp.

Macrosiphum rosae iharae Matsumura- 1

Macrosiphum liriodendri Monelb (Present study)

Megoura viciae japonica (Matsumura)

Melanocallis caryaefoliae (Davis )^ (Present study)

Monellia caryella (Fitch)*- (Present study)

Moneltinpsis pecanis Bissell 2 (Present study)

Myzus malisucta Matsumura 3

Myzus persicae (Sulzerr (Hodek 1973)

Neophyllaphis podocarpi Takahashr* 3 (Present study)

Nippolachnus piri Matsumura^

Periphyllus califarniensis (Shinji) 3

Rhodobium porosum (Sanderson) (Present study)

Rhopaiosiphum pseudobrassicae Davis-

Rhopatosiphum prunifoliae Shinji-

Schizaphis graminum (Rondani) (USDA, APHIS, Niles, MI)

Tinocallis kaiiawaluokulani (Kirkaldy)^ (Chapin & Brou 1991)

Toxoptera odiriae (Van der Coot) 4

Toxoptera piricoia Matsumura^

242 ENTOMOLOGICAL NEWS

Table 5. Recorded prey of Harmonia axyrkiis Pallas (continued)
Taxon and Species fSource(s)] (continued r

Diaspididae

Pseudaulacaspis pentagona (Targioni-Tozzetti) (Park and Kim 1990)

Lepidosaphes salicina Borchsenius 4
Eriocoecidae

RhizoCQCCUS transversus (Green K
Margarodidae

Icerya purchasi Maskelh

Matsucoccus resincmie Bean & Godwin- (McCIure 1987)

Matsucoccus matsumurue (Kuwana) (Kao & Yun 1983r
Phylloxeridae

Phylloxera notabiUs Pergande^ (Present study)
Pseudococcidae

Nesticoccus sinensis Tang-

Phenacoccus pergandei Cockerel 1
Psyllidae

Anomoneura mart Schwarz^

Thysanogyne limhaia Enddeyein 4
Lepidoptera:
Arctiidae

Hyphaniria cunea (Druryh

' Unless indicated otherwise, listed in Yasumatsu and Watanabe (1964) (citing other sources) as
prey species in Japan.

2 Prey species recorded in New World.

3 listed in Chapin and Brou (1991) and citing other sources. Many other synonymous names are
also given but these are not reproduced here,

4 listed in Yarn-/ a/.( 1989).

5 Shu and Yu (1985).

ACKNOWLEDGMENTS

We thank Judy Duff, Wanda Bragg, Faye Brown, Sherrie Yarbrough and Judy Friedel
for their technical assistance during this study. Joseph Tropp reared all beetles at the Newark,
DE, facility. We also thank Billy Ree, Allan Knutson, and two anonymous reviewers for their
suggestions.

Vol, 105, No. 4, September & October, 1994 243

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